The Glasgow Naturalist (online 2017) Volume 26, part 4, xx-xx

Natural history contributions of the University of Glasgow Exploration Society to Scotland and the World

J. Roger Downie1, E. Geoffrey Hancock2, Stewart A. White1, Annette C. Broderick3, Brendan J. Godley3

1 School of Life Sciences, Graham Kerr Building, University of Glasgow, Glasgow G12 8QQ 2 The Hunterian Zoology Museum, University of Glasgow, Glasgow G12 8QQ 3 Centre for Ecology and Conservation, University of Exeter, Penryn Campus, Cornwall TR10 9FE

E-mail: [email protected]

ABSTRACT source that began around the same time was the Expeditions with a natural history focus have been Royal Geographical Society’s expedition grant organised by University of Glasgow staff and scheme (1956). students since the 1930s. The educational benefits of such expeditions to students have been reported by We have not been able to locate any documents Harper et al. (Journal of Biological Education 51, 3- relating to the foundation of the University of 16; 2017). Here, we present a short history of these Glasgow’s Exploration Society (GUExSoc), but the expeditions, concentrating on their scientific first expedition reports held by the University achievements. In addition to expedition reports, a Library which acknowledge the existence and large number of PhD theses, masters and honours support of GUExSoc are for Finland (1970) and project reports and scientific papers have been Calabria (1970), and we were told that Morocco based on expedition work. Many biological (1970) was also organised through GUExSoc specimens have been deposited in museums, (Hansell, pers.comm.). including some new species. We provide case histories of four expedition locations, to demonstrate Some of the expeditions in the late 1960s to early the variety of work done, and the value of returning 1980s were essentially field research trips for the many times to the same place: Scotland, Trinidad and final year class in Topographic Science, aimed at Tobago, North Cyprus and Ecuador. A major problem gathering data for final year dissertations. Although for expeditions is funding. For many years, the the Carnegie Trust initially supported these under Carnegie Trust for the Universities of Scotland ran a the University Expeditions scheme, they later took funding stream that was crucial to the viability of the view that Universities should fund dissertation Scottish university expeditions, but this has sadly work themselves. Later Topographic Science now closed. For Glasgow University expeditions, the expeditions such as Maam, 1978 and Inchnadamph, Blodwen Lloyd Binns Bequest has provided a reliable 1979 (Table 1) therefore received no Carnegie source since 1994, and we hope that it will continue funding. to do so. Membership and equipment lists and accounts INTRODUCTION survive for the early years of GUExSoc, 1972-1982. The first University of Glasgow staff-student Membership averaged about 30 per year and cost expeditions with a natural history focus that we £0.25. The Society had an equipment store mainly know of were to the inner Hebridean island of Canna comprising tents and other camping equipment. in 1936 and 1937 (Table 1A). After a hiatus of a Excluding the Topographic Science field trips, an dozen years, partly explained by the Second World average of two expeditions per year occurred over War, the next such expedition was to the Garvellach that period. Islands in 1949 and then to St. Kilda in 1956. Thereafter, one or two expeditions took place most In 1988 after a few years of inactivity, GUExSoc was years until the early 1980s (Table 1). Initially, these re-constituted with the full support of the University expeditions were organised by interested groups of Principal at that time, Sir Alwyn Williams, and the staff and students, who applied to the University University Court, and, crucially, with a new funding Court for approval and financial support, and to procedure. The 1988 settlement established a new external sources, especially the Carnegie Trust for body, the Exploration Council, technically a sub- the Universities of Scotland (hereafter, Carnegie committee of Court and composed of a small number Trust). In 1959, the Carnegie Trust instituted a of senior staff with relevant interests. As first chair of specific funding stream for University Expeditions the Council, Ian Thomson, Professor of Geography, (Walker, pers. comm.). Another important funding steered this new arrangement through its initial years.

Year Location Themes and Outcomes A) Scotland 1936, 1937 Canna (Inner Hebrides) + Geology, ecology, zoology. Published papers on natural history of Canna and Sanday (Bertram, 1939), birds of Canna (Carrick & Waterston, 1939), and parasites of birds and mammals (Carrick, 1939). Also a report on expedition provisioning (Blair, 1936) 1949 Garvelloch Isles Geology, zoology. Six published papers on marine (Inner Hebrides) lamellibranchs (Hunter, 1951), geology (Hunter & Muir, 1954), insects (Muir, 1954), birds (Dunn et al., 1954), ecology (Hunter, 1954) and molluscs (Hunter, 1953) 1956 St. Kilda # Zoology (two papers: Hamilton (1963), Fraser et al.,1957) (Outer Hebrides) 1962, 1964 South Uist + Zoology. Insect specimens collected (Outer Hebrides) 1974 Callanish, Lewis * Geography (TS) (Outer Hebrides) 1975, 1976 Durness * #Zoology, Botany. The full report for 1975 exists; for 1976, (Sutherland) only a hand-written plant list 1976 Scourie (Sutherland) *Geography (TS) 1979 Inchnadamph * Geography (TS) (Sutherland) 1980 South Uist ~ Carnegie funded; no other record (Outer Hebrides) 1980, 1982 Foula # Bird ringing and other data collection (several papers built (Shetland) on data collected, notably Furness, 1981; Furness Todd, 1984

B) Beyond Scotland 1963 Narvik (Norway) *Geography 1964 La Causse du Larzac *Geography (France) 1965 Austria *Geography 1965, 1966 Iceland Geography. Glacier mapping 1966 Portugal + Zoology (fish, entomology); book chapter by Miller (1986) 1966 Yugoslavia * Geography 1969 Norway * Geography (TS) 1970 Morocco # Zoology 1970 Finland # Geography (TS) 1970 Calabria (southwest Italy) * Geography (TS) 1971 Valais (Switzerland) * Geography 1972 Iceland * Zoology and Geology 1973 Grimsel Pass (Switzerland) Geography (glacier mapping) 1973 Morocco ~ Carnegie funded; no other record 1974 Finland * Geography 1975 Czechoslovakia ~Carnegie funded; no other record 1975 Pyrenees ~Carnegie funded; no other record 1977 Pyrenees ~Talk given and Carnegie funded 1977 Seychelles Zoology. Report may be lost. Two research papers on fairy terns (Houston,1979), and skinks (Brooke & Houston, 1983) 1978, 1980 Maam (Finnish Lapland) * Geography (TS) 1978 Corsica * Geography 1979 Faroe Islands ~Carnegie funded; no other record 1980-81 Crete ~Carnegie funded; no other record 1982 Cwm Idwal (Wales) * Geography (TS) 1983 Newborough Warren * Geography (TS) (Wales)

Table 1. University of Glasgow Expeditions 1936-1983. Usage of * denotes report in University of Glasgow Library; + denotes specimens deposited in the Hunterian Museum; TS denotes Topographic science class field trips; ~ denotes expeditions known only from the records of the Carnegie Trust for the Universities of Scotland or from GUExSoc minutes books.

Year Number and Years Themes a) Scotland Islay (inner Three: 2014-16 + Zoology Hebrides)

b) Europe Azores Two: 1989, 2003 Ornithology North Cyprus Eight: 1992-98 + Marine turtle conservation/biology; feral donkeys Cyprus Fourteen: 2000-16 Marine turtle conservation/biology (RAF Akrotiri) Iceland Eighteen: 1992-2002 First series: glaciology; second series: wildlife and 2008-16 biology/conservation Spitzbergen 1996 only Geography

c) Africa Canary Islands Two: 2000, 2016 Geology Egypt Five: 1990; 2013-16 First, mainly geography, plant ecology; later series ecology Gambia Four: 2005-08 welfare, zoology Kenya Two: 1998, 2016 First, zoology; more recent, sustainable development Seychelles 1996 only Zoology Tanzania Eight; 1991, 1998, Mainly geographical, but also zoology 2008-15 Zambia Two; 2006, 2008 Aquatic plant ecology, zoology

d) Asia Borneo 1999 only Zoology Himalayas 1993 only Geography Kazakhstan 1998 only Ornithology Oman 1990 only Ornithology

e) Australasia Kimberley 1996 only Geography (Australia) Tutamoe 1995 only Geography (New Zealand)

f) North America Canada 1997 only Wildlife conservation

g) South America and the Caribbean Bolivia Eleven: 1998-2016 + Zoology, ornithology Brazil Three: 1999-2003 Aquatic plants, zoology Costa Rica 2005 only Zoology Ecuador Twelve: 1997; + Zoology, ecotourism development 2000-2012 Guyana 1995 only + Zoology Peru Seven: 2006-2015 Zoology Trinidad and Thirty five + Zoology, some plant ecology, geology Tobago expeditions: 1989-2016: two to T and T; 20 to Trinidad alone; 13 to Tobago alone

Table 2. University of Glasgow Expeditions 1989-2016. Usage of + denotes specimens deposited in the Hunterian Museum; reports for most expeditions are available in the University Library and/or on the GUExSoc web-site; we have not attempted to collate the large number of publications resulting from these expeditions; see the text, especially the case histories.

Council was given an annual budget and the task of projects, including expeditions (e.g., Trinidad 2000 overseeing the activities of GUExSoc, especially of in TGN 23(6), 2001; Azores 2003 in 24(2) 2004). scrutinising expedition proposals, deciding whether More recently, summary expedition reports have or not to approve them, and then deciding on the appeared in the GNHS quarterly newsletter and full level of funding to provide. GUExSoc itself began reports have been accessible via GUExSoc’s web-site afresh in 1988 with a new constitution. The first of (http://glasgowexsoc.org.uk/). the new era of expeditions took place in 1989 (Table 2: Trinidad and Tobago; Azores). The constitution The educational value to students of participating in made clear that the main aim of expeditions should expeditions has been reported by Downie et al. be the advancement of science; approval and support (2008), describing the potential of expeditions for would not be given to proposals that simply involved students undertaking final year research projects, adventurous travel. and by Harper et al. (2017) who emphasise the transferable skills/graduate attainment value of The conditions of funding from the Carnegie Trust organising and taking part in expeditions. In this were important to all Scottish Universities which paper, we focus on the natural history research organised expeditions. The Trust’s rules were that outcomes of the long series of expeditions run by expeditions should mainly be for undergraduate GUExSoc. Given the very large number of expeditions students with staff providing support and training; organised through GUExSoc, it would be impossible that the sponsoring University should provide to do justice to them all. We therefore have adopted formal approval of each expedition proposal and a case-history approach and present four themes: some level of financial support; and that work aimed Scotland, Trinidad and Tobago, North Cyprus, and at student dissertations should not be the main Ecuador. purpose of an expedition. Correspondence with the Trust clarified that individual students were not EXPEDITION CASE HISTORIES barred from carrying out dissertation research as Scotland long as this was not the focus of the expedition as a All the earliest expeditions were to Scottish whole; also that full-time staff presence was not localities, some visited more than once, from the essential. In the early years, Trust grants were a few earliest to Canna in 1936 up to Foula in 1980 and hundred pounds per expedition; when the scheme 1982 (Table 1A). The first location, Canna plus ended in 2014, the maximum award was £2K per neighbouring Sanday, was visited in consecutive expedition. Of the seven Scottish Universities which years and produced high quality results published in benefitted from Trust expedition grants, GUExSoc peer-reviewed journals (Bertram, 1939; Carrick, was the most active (Table 3). 1939; Carrick & Waterston, 1939). These described many aspects of the natural history of the islands: Glasgow Natural History Society (GNHS) has had an physical geography, botany, terrestrial and important role in GUExSoc expeditions following the freshwater invertebrates, birds and mammals. The establishment of the Blodwen Lloyd Binns Bequest impressive range of coverage was the product of the (BLB) in 1993. The BLB committee agreed in 1994 students plus a considerable number of experts who that undergraduate expeditions were one of the helped identify pre-sorted specimens. Bertram activity categories that would be eligible for support. (1939) wrote that ‘The original object was to survey In the period 1994-2010, the Bequest supported 59 an island in the West of Scotland with a view to expeditions, granting a total of £33.5K, 22% of its providing data which would serve as a basis for annual grant expenditure (Downie et al., 2012), and future ecological studies’. Sadly, the Second World a similar level of support (£5K per annum) has War got in the way of the planned future work. The continued since then. In addition, GNHS has hosted standard of the directly published outputs from the many talks on these expeditions in its talks two Canna expeditions has never been exceeded. programme, and for a few years, The Glasgow Naturalist included short reports on BLB funded

University Number of Total awards Percentage of total grants (£: rounded awards thousands) St. Andrews 29 £16K 2.9 Edinburgh 106 £125K 23.6 Glasgow 160 £243K 45.9 Aberdeen 83 £77K 14.6 Dundee 25 £29K 5.6 Stirling 15 £12K 2.4 Napier 13 £26K 4.9 Joint SUs 1 £.5K 0.1 Table 3. University expedition grants awarded by the Carnegie Trust for the Universities of Scotland, 1959-2014. Data kindly provided by Professor Andy Walker, Secretary and Treasurer of the Trust.

Twelve students were involved over the two years, The two expeditions to the Shetland island of Foula mainly zoologists but including two botanists. The (1980, 1982) were exceptional, firstly in having expedition’s logistics also were published as a guide participants from several other organisations. Of the to future similar endeavours. Blair (1936) noted that 11 members of Foula (1982), six were Glasgow based an investigation into dietary needs on expeditions and the others from Durham, Edinburgh, would have been of interest but that “one could not Peterborough and London. Secondly, these risk impairing the members’ efficiency by expeditions did have published outputs (Table 1A). experiments in another sphere of science”. In addition, Furness (pers. comm.) reports that these expeditions collected baseline data that contributed The next expedition was to the Garvellach Isles to several later papers, notably an important paper (1949). This also led to an impressive list of six in Nature by Votier et al.(2004), demonstrating the papers, after some delay (Table 1A). One of the huge value of long-term datasets in ecology. students and authors, David A. Muir, who went on to a career in malaria research, has provided some After a gap of three decades, in response to the needs reminiscences of the expedition, which had the of students who could not afford the expense of grand title: Scottish Ecological Expedition to the overseas expeditions, and also who planned careers Garvelloch Islands (SEEGI). Note that the name of in UK conservation and environmental management, these islands in English is a version of the Gaelic GUExSoc introduced the idea of a ‘Remote Scotland’ Garbh Eileaich; in the 1940s this was given as expedition in 2013. This clearly met a need, as many Garvelloch, but more recently as Garvellach; we use students applied, even though the location was the latter spelling here, except for the titles of the initially undecided. Islay was soon chosen (not, original publications. There are some butterfly perhaps, particularly ‘remote’) partly because of the specimens in the Hunterian Museum with these input of the RSPB and also because of apparently enigmatic initials on their labels (Muir, correspondence with the Islay Natural History Trust 1954, is a short note on the Garvellach insects). The which was enthusiastic about the idea. The 2017 students were accompanied by staff member Bill Islay expedition was the fourth in succession. In Russell Hunter, who later had a distinguished career addition to the normal reports, the expeditions have in the USA. Muir provides a piece of doggerel helped add to the biological records of the island commenting on Hunter’s cooking skills: (Fig.1).

‘Poor Hunter, marooned with his mutinous crew And all because of his pemmican stew! He cooked it in secret with weird incantations And added some nettles to spin out the rations. He served it up steaming and reeking and rich, Dark golden in colour and sticky as pitch. But of course all the ‘gannets’ with palates of leather Simply couldn’t detect the delectable flavour! They complained it was gritty and cooked in a hurry And proceeded to drown it in Worcester and curry. By this time the Dixie was only half emptied Fig. 1. Leaders Lorna Archer and Angus Lothian checking But the ‘gannets’ shied clear and refused to be wildflower identification, Islay 2015 (photo credit: tempted Richard Thompson) For only a staunch and redoubtable few Would dare a repeat of that pemmican stew. Trinidad and Tobago Yes, Hunter concocted many a brew, When GUExSoc was re-constituted in 1988, one of But never a one like the pemmican stew!’ the first expeditions was to Trinidad and Tobago in 1989. Why there? One of us (JRD) had already St. Kilda (1956) also had published results (Table established a research link in Trinidad and was 1A) but thereafter the main outcomes were aware of the potential of the islands as a location for expedition reports, descriptive of the projects a student expedition. In addition to its under- undertaken. An important purpose of these was to researched biodiversity, the country was peaceful, provide funders with prompt accounts of the work relatively small and therefore accessible, English- achieved, and they remain the only account of most speaking and mostly free of troublesome tropical of these activities. The practice of depositing a copy diseases. In Trinidad, also, the University of the West in the University Library began in the 1960s but was Indies (UWI) campus at St. Augustine offered a base not always adhered to (Tables 1 and 2). with laboratory facilities, should we need them. Even with these advantages, it is unlikely that any of the 1989 participants envisaged that they would be the

first of 35 expeditions spanning (by the end of 2016) working together in flexible teams. Table 4 shows the 27 years (Table 2). distribution of project themes.

Looking back, the 1989 expedition was almost Work on and marine turtles has dominated, recklessly ambitious. The total personnel list reflecting the research interests of the staff members included 33 people, a mix of staff, post-graduates and mainly involved. Other themes have often resulted undergraduates, led by Richard Rutnagur, a PhD from interactions with local organisations: for student with Trinidadian family connections. The example, studies on monkeys and bats arose from expedition worked on both islands with the Trinidad requests by the Trinidad Government’s Wildlife team based near UWI, and the Tobago team split Section, and marine turtle monitoring on Tobago between two locations; marine biologists in the from 2004 resulted from interactions with local south at Buccoo and botanists in the north at NGOs, Save Our Sea Turtles (SOS) and North East Sea Charlotteville. Funding an expedition of this size was Turtles (NEST). Each expedition has produced a difficult and co-ordinating activities in the days report, available in the Glasgow University Library, before mobile phones (in a country with a barely with most also available on GUExSoc’s web-site functioning telephone system) was more so. Despite (www address: https://glasgowexsoc.org.uk ). In these problems, the expedition produced a report of addition, five PhD students have carried out their nearly 100 pages, covering the results of a very field research in association with Trinidad diverse set of projects: the state of Tobago’s coral expeditions, and several Masters students and many reefs; regeneration of the Tobago Main Ridge forest undergraduates have undertaken their research following the devastation caused by Hurricane Flora projects on the islands. All this research has in 1965; the distribution and habits of an endangered contributed to over 90 scientific papers in peer- endemic bird, the Trinidad piping guan; the nesting reviewed journals. In addition, the work has activities of leatherback turtles on Trinidad’s east contributed to a recent field guide to the reptiles and and north coast beaches; conservation measures of Trinidad and Tobago (Murphy et al., suitable to protect the collared peccary whose 2017) and to a book chapter on the conservation populations are heavily affected by hunting; and status of the islands’ frogs (Auguste et al., 2017). several projects on the reproductive ecology of Some might think this a surprising outcome for work Trinidad’s frogs. mainly done by undergraduate students, but if such work is properly supervised, there is no reason why The second expedition (1991) was similar, also with it should not produce competent and publishable 33 participants and with work on both islands, science, especially from locations holding such an generating a report of 123 pages: in Tobago, only on abundance of biodiversity and where basic natural the coral reefs in the south; in Trinidad, more on history observations are still needed. One of the marine turtles, the piping guan, reproduction pleasing outcomes of the Trinidad and Tobago and new projects on dragonfly territoriality and expeditions is how often they have helped young veterinary aspects of public health, involving scientists to achieve their first publications. Of the vultures and goats. 90+ papers derived from these expeditions, over 60 of the authors were undergraduates when they did Thereafter, expedition teams were smaller and the work (see Downie, 2012 for a complete list of the based on only one of the islands: 20 more publications to 2010). In addition, many Trinidad expeditions to Trinidad (1993-2016, almost yearly) and Tobago expedition participants have progressed and 13 consecutive visits to Tobago (2004-16). Most to distinguished careers in science. What has all this of the expeditions have included several distinct effort achieved? We present a short selection of projects (five on average) with staff and students highlights.

Themes 51.1 Frogs: reproductive ecology, behaviour, treefrog adhesion, distribution, new species, phylogeny, conservation 20.9 Marine turtles: nest monitoring and conservation, nesting behaviour and physiology, hatchling behaviour 13.2 Invertebrates: crab behaviour, insect systematics and ecology, corals- fossil reefs and reef conservation 7.1 Birds: diversity, behaviour, ecology 4.9 Mammals: wild pig conservation, monkey behaviour, bat diversity 1.6 Lizards: diversity and behaviour 1.1 Fish: diversity, ecology

Table 4. Distribution of Trinidad and Tobago expedition project themes, from 35 expeditions and 182 projects.

The golden tree frog Phytotriades trinitatis (Fig.2) is government and local communities that led to the one of Trinidad’s most iconic species. Living in the establishment of a set of NGOs whose role was to water tanks enclosed by the leaves of the giant protect nesting turtles and to educate visitors about bromeliad erectiflora, this frog is them. Our later expeditions have worked with found only near the summits of the island’s highest several of these NGOs: at Matura, Grande Riviere, mountains. Using molecular phylogenetic methods, Matelot and Fishing Pond (in Trinidad) and with SOS Jowers et al. (2008) established that this species and NEST in Tobago. Our work has helped clarify the should be classed in a genus on its own and that it is sizes of the nesting populations and to map the not closely related to frogs in the genus beaches where they occur (Walker et al., 2015). to which it was previously assigned. More recently, Brozio et al. (2017) have established the value of the new method of environmental DNA for detecting these frogs without having to destroy their host plants. Monitoring the health of the golden tree frog population should be much easier in future.

Fig. 2. The golden tree frog, Phytotriades auratus, El Tucuche, Trinidad (photo credit: Gillian Simpson)

Before our work, the stream frog Mannophryne trinitatis was considered to occur both in Trinidad Fig. 3. A male Trinidad stream frog, Mannophryne trinitatis and in the nearby Paria Peninsula of Venezuela, and transporting his tadpoles (photo credit: Joanna Smith) possibly also in Tobago. Manzanilla et al. (2007) showed that the Venezuelan and Tobagonian populations belong to different species, confirming M. trinitatis as a Trinidad endemic (and the Tobago species M. olmonae as a Tobago endemic). In Mannophryne species, males guard the eggs on land, then carry the hatched tadpoles to water (Fig.3). Downie et al. (2001) showed that transporting males are careful to avoid depositing their tadpoles in water containing predators and that they can carry their tadpoles for several days in search for a suitable pool or stream. An early assessment by IUCN suggested that this species is amongst the frog species threatened with extinction, but our recent results (Greener et al., 2017) show that it is highly abundant and widespread in suitable habitat in Trinidad. Fig. 4. Entomologist Jeanne Robinson sampling from a bromeliad, Arima Valley, Trinidad (photo credit: Geoff The first two Trinidad expeditions included Hancock) monitoring of the east and north coast beaches for Several new species of Diptera (flies) have been nesting marine turtles, at the request of the Wildlife named and described as a result of our Trinidad Section’s director, Dr Carol James. The findings from expeditions. The phorid Megaselia nidanurae was these expeditions (later written up by Godley et al., discovered to lay its eggs in the foam nests of the 2001a, b) contributed to the discussions between burrow-nesting frog Leptodactylus fuscus (Downie et

al., 1995). A new species in the Mycetobiidae, and BJG were awarded University of Glasgow post- Mycetobia downiei was found as larvae in sap runs of graduate scholarships to conduct the fieldwork for mature Saman and Mora trees (Hancock, 2016). their PhDs in Cyprus. This amazing opportunity Arising from observations made in bromeliads of fly secured the long-term future of the project and larvae while monitoring the golden tree frog (Fig.4), formed a platform for both academic careers. some projects were planned. Hoverflies were reared from bromeliads and similar phytotelmatus habitat and three new species Copestylum elizabethae, C. louisae (Rotheray et al., 2007) and Quichuana longicauda (Ricarte et al., 2012) were revealed.

North Cyprus In 1992, the first Glasgow University Turtle Conservation Expedition (GUTCE) to Northern Cyprus took place, in many ways resulting from the Trinidad and Tobago expeditions of 1989 and 1991. Plans to study wild birds on the 1989 Trinidad and Tobago expedition were abandoned at the last minute due to permit issues and instead it was suggested that the group work on marine turtles.

Brendan Godley (BJG) was part of the 1989 team and Fig. 5. The turtle team at The Goatshed, main base of the returned (with ACB) as leader in 1991 to continue marine turtle project, North Cyprus (photo credit: Annette the work on marine turtles. Meanwhile, a fortuitous Broderick) meeting took place in Cyprus in the summer of 1991, between Kutlay Keço, Ian Bell, and Celia Moorley- Although mostly focusing on marine turtles, other Bell of the Society for the Protection of Turtles in studies have been conducted on crabs, frogs, birds, Northern Cyprus (SPOT), and a friend of University dune vegetation and a separate GU expedition was of Glasgow staff members Sally Solomon and Roger mounted to study the feral donkeys. The project has Tippett. The need to do more to protect the turtles in continued each year since and is now in its 25th year, Cyprus was discussed, and aware that Sally had although it subsequently moved with BJG and ACB to conducted research on turtles, the friend suggested the University of Swansea (1999) and then Exeter that the SPOT team write to Sally to ask if there was (2003). Now named the Marine Turtle Conservation a group of students who might come out to Cyprus to Project, it is a collaboration with the Society for the help. Knowing that BJG had been involved in Protection of Turtles in Northern Cyprus (SPOT) and monitoring marine turtles in Trinidad and Tobago, the Environment Protection Department and accepts Sally passed on the letter and asked BJG if he could volunteers from all around the world. The Goatshed get a group of students together to go out to Cyprus is still our main base on the island and also now has in the summer of 1992. The project was born and a a small information centre for visitors and local group of 14 students, led by BJG and ACB and people to come to find out about turtles and if supported by University of Glasgow staff members possible arrange to see nesting and/or hatchling including Sally Solomon, Roger Tippett and JRD took turtles. part in the first GUTCE expedition. Overall the project has led, in total or in part, to eight In 1992, our main base was at the Fire Station in PhD theses, with a further five underway, and has Yesilkoy on the Karpaz peninsula, and as we resulted in over fifty peer-reviewed scientific papers. explored all of the 80+ turtle nesting beaches over In addition, data from the project have provided the the season, we quickly came to realise the basis for the designation of five Special importance of Alagadi Beach, situated on the north Environmental Protected Areas (SEPAs) containing coast of the island, which was also conveniently turtle nesting beaches. These sites have also been where SPOT President Kutlay Keço lived. In that first identified as potential Natura 2000 sites and are year we camped on the beach, which was pretty awaiting designation by the European Union. The tough after a night shift, trying to sleep in the heat of duration of the project means that over 1200 the day! Over that summer however, Ian and Keço students have been able to experience a summer of came to the conclusion that we were a hard- working challenging ecological research, environmental and eager group and asked us to return the following education and direct conservation action, with many year, generously renovating an old building at moving on to careers in these sectors. The student Alagadi for us, named the ‘Goatshed’ (Fig.5) after the group has always had an international feel with previous inhabitants! During the summer of 1993 students from across the world partaking. In recent we came to realise that there was a wealth of years, growing numbers of Cypriot students studying research to be conducted on the turtle populations in veterinary and life sciences have been working as Cyprus and thanks to Sally’s persistence, both ACB part of the project which augments the effectiveness

of the project and the cultural experience for all interviewing over 40 applicants a group was involved. selected. The first Ecuador expedition was comprised of two research groups, a bird group led A full list of publications can be found at by Stewart White and an insect group led by GH from but a highlight has been the the Hunterian Museum and Graham Rotheray from satellite tracking studies over the years that have the National Museums of Scotland. Both groups told us so much we didn’t know about the life spent an initial period of three weeks at the private histories of both green (Chelonia mydas) and cloudforest reserve of Otonga in the Andes south of loggerhead (Caretta caretta) turtles. We have Quito. After this initial period of research the bird described their foraging locations - mostly along the group travelled on to Amazonia and met up with north African coast, their over-wintering behaviour - Darwin and his brother Edwin to travel to Payamino. moving offshore to deeper waters in the colder The first year saw many teething problems, months where they do little but breathe – indeed we inevitable when taking a group of students to a recorded the longest dive of an air breathing country for the first time and particularly when only vertebrate – a loggerhead turtle whose dive lasted two of the group had very basic Spanish and no 10.5 hours and made it into the Guinness Book of knowledge of kichwa. The expedition was enough of Records! A second highlight has been that two of our a success, however, to encourage Stewart to arrange PhD students have married Cypriot scientists and with Darwin and Giovanni Onore the owner of they are now living in Cyprus with their families and Otonga to make regular future visits. all working on ecological/environmental issues with NGOs or Universities. At the time of writing there have been 12 University of Glasgow Ecuador expeditions, with the focus of the Funding has always been a challenge, given the visits having gradually moved towards political sensitivities of working in the north of concentrating on Payamino. Well over 100 Glasgow Cyprus, yet small pots of money from numerous undergraduate and postgraduate students have funding bodies and hundreds of self-funded student visited Ecuador on expedition, conducting research volunteers who have worked day and night over the on leafcutter ants, butterflies, dragon and years have enabled the project to continue annually damselflies, fish, frogs and mammals. The bird for more than two decades. We thank them all for research has continued to be the main project with their support. the species list for Payamino now standing at 340 species and with several thousand individually Sadly, Ian Bell and Sally Solomon are no longer with marked birds flying around the rainforest carrying us but the legacy of their generous mentorship and Swedish aluminium bands marked ’University of support continues. Glasgow’, plus a few marked ‘University of Dundee’, but’s that’s another story! Ecuador A one-off expedition to northeast Ecuador took place As well as expeditions, an undergraduate field in 1997, working on bird populations in degraded course, part of the final year Tropical Rainforest forest, and on begonias. However, a later series of Ecology option has been running for the last 11 University of Glasgow expeditions to Ecuador came years. This gives the students the chance to study the about due to one of those happy coincidences that rainforest ecosystem first hand and conduct short happen only very occasionally. Glasgow Zoology group research projects. The students selected have graduate Nan Swannie (a member of the Trinidad to pay a personal contribution but the field course is expedition in 1993) visited Ecuador on holiday and heavily subsidised by the University – evidence that sampled some ecotourist experiences. By chance, the value of field teaching in such a biodiverse wandering along the road in the Amazonian oil town environment is appreciated. of Coca she met a tour guide called Darwin Garcia and took his tour up the Rio Payamino to the The expeditions and field courses have given the community of San José de Payamino. Not long after students involved a unique experience and several she returned home, Nan visited Glasgow to give a talk who visited as ‘ordinary’ expedition members later on her experiences in Ecuador to the student returned as expedition leaders and in two cases as Zoological Society. At the end of her presentation she members of staff on University field courses in mentioned that Darwin was keen to have scientific Payamino. Several of the students have gone on to research conducted in the Payamino area. SW was in post-graduate study and all would say that the the audience and already a veteran of multiple Ecuador Expedition or field course was the most student expeditions to Trinidad. He chatted to Nan memorable part of their time at University. Along over the usual post-lecture cheese and wine and, with the University of Manchester and Universidad despite already having committed to a Trinidad Estatal Amazonico in Puyo we have been able to expedition, agreed to propose an Ecuador expedition found a permanent research station at Timburi for the following summer, 2000. Cocha on the bank of the Rio Payamino (Fig.6). This The Ecuador Expedition proposal went down well now welcomes researchers from Ecuador, North with students in Exploration Society and after America and Europe and has seen the publication of

several papers in peer-reviewed journals (e.g. Muir children to learn about their local wildlife. Many & Muir, 2011; White & Patino, 2017), with more in other examples could be given; for more information, preparation. From entomological work during two of read the expedition reports in the GUExSoc archive. the Otonga cloud forest expeditions a number of hoverflies (Syrphidae) were reared from bromeliads DISCUSSION and decaying plant material of two new species, It is clear from the account provided here that Copestylum ontongaensis and C. tapiai (Rotheray et University of Glasgow expeditions have generated a al., 2007). All this was the result of a chance meeting vast amount of natural history information, both in in the dusty streets of Coca. Scotland and abroad, and that the size and scope of the expedition enterprise has grown hugely since the 1930s. It is also clear from this account and the research by Harper et al. (2017) that the expedition experience has been of great benefit to young people starting their careers in science. To finish this historical review, we ask some questions on sustainability and future aims. The loss of Carnegie funding has been very damaging to expeditions organised by Scottish Universities, and we hope that in time, the Trust will re-consider and develop a new scheme. Despite the demonstrable value of expeditions, the University of Glasgow itself does not have a clearly ear-marked method for funding them. We hope that this can be resolved in the near future. The Blodwen Lloyd Binns Bequest has been a valuable and reliable source of grants, but it does not Fig. 6. Glasgow students and local people interacting at the possess the resources to be a major funder. As for the Rio Payamino community, Ecuador (photo credit: Stewart student population, the large number who attend the White). annual proposals meeting, where the next summer’s expeditions are outlined, and students are invited to EDUCATION AND OUTREACH apply for places, shows no signs of diminishing Expeditions are educational in many ways: the enthusiasm. Students recognise the value of an participants learn a lot about the places they visit, opportunity to carry out real research which they develop new skills and learn about their own have helped plan themselves, and which is carried capabilities, as discussed by Harper et al. (2017). The out in a location far from their previous experience. results presented in the many publications derived However, although the students raise some of the from expeditions provide information on the places funds through their own activities and personal visited. However, expeditions have additional contributions, the ever-rising costs of expeditions educational roles. In most cases, they work in indicate the need for reliable external funding. As we collaboration with local organisations, sometimes have shown, Glasgow expeditions have been to government agencies, more often NGOs, as outlined locations both in Scotland and abroad. Both have in the earlier case histories. These collaborations generated valuable results and experiences, and we provide many opportunities for two-way learning. In hope that they can continue into the future. Although addition, many expeditions have built into their aims Scottish based expeditions are less expensive and explicit educational activities. In the marine turtle easier to organise, Scotland has a long history of focused expeditions run in both North Cyprus and at engagement with other countries, and it is still the the RAF base at Akrotiri in the south, educating case that in many countries, the experiences, skills beach visitors about the turtles, threats to them and and interests of Scottish researchers and students how to avoid causing harm have been constant can make valuable contributions to the expansion of activities. For several years, the Trinidad expeditions knowledge. were involved in a British Council sponsored link between Glasgow and Trinidad schools; this ACKNOWLEDGEMENTS involved working in Glasgow schools, carrying out We thank David A. 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